Marcello A. Mannino

The genetic history of Ice Age Europe

Modern humans arrived in Europe ~45,000 years ago, but little is known about their genetic composition before the start of farming ~8,500 years ago. We analyze genome-wide data from 51 Eurasians from ~45,000-7,000 years ago. Over this time, the proportion of Neanderthal DNA decreased from 3–6% to around 2%, consistent with natural selection against Neanderthal variants in modern humans. Whereas the earliest modern humans in Europe did not contribute substantially to present-day Europeans, all individuals between ~37,000 and ~14,000 years ago descended from a single founder population which forms part of the ancestry of present-day Europeans. A ~35,000 year old individual from northwest Europe represents an early branch of this founder population which was then displaced across a broad region, before reappearing in southwest Europe during the Ice Age ~19,000 years ago. During the major warming period after ~14,000 years ago, a new genetic component related to present-day Near Easterners appears in Europe. These results document how population turnover and migration have been recurring themes of European pre-history.

Sampling shells for seasonality: oxygen isotope analysis on shell carbonates of the inter-tidal gastropod Monodonta lineata (da Costa) from populations across its modern range and from a Mesolithic site in southern Britain

The degree of resolution of seasonal estimates using stable oxygen isotopes from marine shells is determined by various factors, including the potential of the species selected to track annual shifts of temperatures and the size of the annual growth increments. Where the amount of annual shell growth is small, seasonal resolution of samples of shell material might be affected by time averaging of isotopic signals. There is also a need to control the number of samples analysed to affordable levels, which implies planning of appropriate sampling strategies. In this study, we investigate variations in δ18O in both modern and archaeological specimens of the inter-tidal gastropod Monodonta lineata (da Costa), in long sequences of samples (across more than one full year of growth) and in short sequences (three samples) taken back from the edge of the shell. Studies on a number of living populations across the modern range of the species, and at two selected localities over a full year, showed that the δ18O composition of the growing shells generally corresponds to seasonal temperature change. Applying these approaches to shells of M. lineata from the Mesolithic site of Culverwell, Isle of Portland, it was shown that season of death could be demonstrated by interpreting both patterns of edge sequences and actual edge values of δ18O. For this site, season of death was shown to range mainly from autumn to winter, with the possibility of some early spring collection. For Mesolithic communities relying heavily on shellfish and probably other marine and coastal resources, the identification of this season of death has direct bearing on their scheduling of subsistence behaviour and on patterns of settlement.

The makers of the Protoaurignacian and implications for Neandertal extinction

Cultural prehistory in southern Europe The Protoaurignacian culture appeared in the southern European archeological record around 42,000 years ago and was characterized by artefacts including personal ornaments and bladelets. Archaeologists have debated whether it was ancestral Homo sapiens or Neandertals who made these tools and ornaments. Benazzi et al. analyzed dental remains from two Protoaurignacian sites in Italy and confirm that they were H. sapiens. The arrival of this culture may have led to the demise of Neandertals in these areas (see the Perspective by Conard et al.). Science, this issue p. 793; see also p. 754 The Protoaurignacian culture in southern Europe involved anatomically modern humans and overlapped in time with the last Neandertals. [Also see Perspective by Conard] The Protoaurignacian culture is pivotal to the debate about the timing of the arrival of modern humans in western Europe and the demise of Neandertals. However, which group is responsible for this culture remains uncertain. We investigated dental remains associated with the Protoaurignacian. The lower deciduous incisor from Riparo Bombrini is modern human, based on its morphology. The upper deciduous incisor from Grotta di Fumane contains ancient mitochondrial DNA of a modern human type. These teeth are the oldest human remains in an Aurignacian-related archaeological context, confirming that by 41,000 calendar years before the present, modern humans bearing Protoaurignacian culture spread into southern Europe. Because the last Neandertals date to 41,030 to 39,260 calendar years before the present, we suggest that the Protoaurignacian triggered the demise of Neandertals in this area.

Origin and diet of the prehistoric hunter-gatherers on the Mediterranean island of Favignana (Ègadi Islands, Sicily)

Hunter-gatherers living in Europe during the transition from the late Pleistocene to the Holocene intensified food acquisition by broadening the range of resources exploited to include marine taxa. However, little is known on the nature of this dietary change in the Mediterranean Basin. A key area to investigate this issue is the archipelago of the Ègadi Islands, most of which were connected to Sicily until the early Holocene. The site of Grotta d’Oriente, on the present-day island of Favignana, was occupied by hunter-gatherers when Postglacial environmental changes were taking place (14,000-7,500 cal BP). Here we present the results of AMS radiocarbon dating, palaeogenetic and isotopic analyses undertaken on skeletal remains of the humans buried at Grotta d’Oriente. Analyses of the mitochondrial hypervariable first region of individual Oriente B, which belongs to the HV-1 haplogroup, suggest for the first time on genetic grounds that humans living in Sicily during the early Holocene could have originated from groups that migrated from the Italian Peninsula around the Last Glacial Maximum. Carbon and nitrogen isotope analyses show that the Upper Palaeolithic and Mesolithic hunter-gatherers of Favignana consumed almost exclusively protein from terrestrial game and that there was only a slight increase in marine food consumption from the late Pleistocene to the early Holocene. This dietary change was similar in scale to that at sites on mainland Sicily and in the rest of the Mediterranean, suggesting that the hunter-gatherers of Grotta d’Oriente did not modify their subsistence strategies specifically to adapt to the progressive isolation of Favignana. The limited development of technologies for intensively exploiting marine resources was probably a consequence both of Mediterranean oligotrophy and of the small effective population size of these increasingly isolated human groups, which made innovation less likely and prevented transmission of fitness-enhancing adaptations.

New chronology for Ksâr ‘Akil (Lebanon) supports Levantine route of modern human dispersal into Europe

Significance Bayesian modeling of AMS radiocarbon dates on the marine mollusk Phorcus turbinatus from Ksâr ‘Akil (Lebanon) indicates that the earliest presence of Upper Paleolithic (UP) modern humans in the Levant predates 45,900 cal B.P. Similarities in early UP lithic technology and material culture suggest population dispersals between the Levant and Europe around 50,000–40,000 cal B.P. Our data confirm the presence of modern humans carrying a UP toolkit in the Levant prior to any known European modern human fossils and allow rejection of recent claims that European UP modern humans predate those in the Levant. This result, in turn, suggests the Levant served as a corridor for the dispersal of modern humans out of Africa and into Eurasia. Modern human dispersal into Europe is thought to have occurred with the start of the Upper Paleolithic around 50,000–40,000 y ago. The Levantine corridor hypothesis suggests that modern humans from Africa spread into Europe via the Levant. Ksâr ‘Akil (Lebanon), with its deeply stratified Initial (IUP) and Early (EUP) Upper Paleolithic sequence containing modern human remains, has played an important part in the debate. The latest chronology for the site, based on AMS radiocarbon dates of shell ornaments, suggests that the appearance of the Levantine IUP is later than the start of the first Upper Paleolithic in Europe, thus questioning the Levantine corridor hypothesis. Here we report a series of AMS radiocarbon dates on the marine gastropod Phorcus turbinatus associated with modern human remains and IUP and EUP stone tools from Ksâr ‘Akil. Our results, supported by an evaluation of individual sample integrity, place the EUP layer containing the skeleton known as “Egbert” between 43,200 and 42,900 cal B.P. and the IUP-associated modern human maxilla known as “Ethelruda” before ∼45,900 cal B.P. This chronology is in line with those of other Levantine IUP and EUP sites and demonstrates that the presence of modern humans associated with Upper Paleolithic toolkits in the Levant predates all modern human fossils from Europe. The age of the IUP-associated Ethelruda fossil is significant for the spread of modern humans carrying the IUP into Europe and suggests a rapid initial colonization of Europe by our species.

Middle Paleolithic and Uluzzian human remains from Fumane Cave, Italy

The site of Fumane Cave (western Lessini Mountains, Italy) contains a stratigraphic sequence spanning the Middle to early Upper Paleolithic. During excavations from 1989 to 2011, four human teeth were unearthed from the Mousterian (Fumane 1, 4, 5) and Uluzzian (Fumane 6) levels of the cave. In this contribution, we provide the first morphological description and morphometric analysis of the dental remains. All of the human remains, except for Fumane 6, are deciduous teeth. Based on metric data (crown and cervical outline analysis, and lateral enamel thickness) and non-metric dental traits (e.g., mid-trigonid crest), Fumane 1 (lower left second deciduous molar) clearly belongs to a Neandertal. For Fumane 4 (upper right central deciduous incisor), the taxonomic attribution is difficult due to heavy incisal wear. Some morphological features observed in Fumane 5 (lower right lateral deciduous incisor), coupled with the large size of the tooth, support Neandertal affinity. Fumane 6, a fragment of a permanent molar, does not show any morphological features useful for taxonomic discrimination. The human teeth from Fumane Cave increase the sample of Italian fossil remains, and emphasize the need to develop new methods to extract meaningful taxonomic information from deciduous and worn teeth.

Climate-driven environmental changes around 8,200 years ago favoured increases in cetacean strandings and Mediterranean hunter-gatherers exploited them.

Cetacean mass strandings occur regularly worldwide, yet the compounded effects of natural and anthropogenic factors often complicate our understanding of these phenomena. Evidence of past stranding episodes may, thus, be essential to establish the potential influence of climate change. Investigations on bones from the site of Grotta dell’Uzzo in North West Sicily (Italy) show that the rapid climate change around 8,200 years ago coincided with increased strandings in the Mediterranean Sea. Stable isotope analyses on collagen from a large sample of remains recovered at this cave indicate that Mesolithic hunter-gatherers relied little on marine resources. A human and a red fox dating to the 8.2-kyr-BP climatic event, however, acquired at least one third of their protein from cetaceans. Numerous carcasses should have been available annually, for at least a decade, to obtain these proportions of meat. Our findings imply that climate-driven environmental changes, caused by global warming, may represent a serious threat to cetaceans in the near future.

Finding the early Neolithic in Aegean Thrace : the use of cores

Using a new approach that combines high-quality coring with AMS dating, the authors are mapping the start of the Neolithic in Aegean Thrace – a missing link in the arrival of agriculture in Europe. The method also revealed the edge of the marine transgression dating to some 2900 years ago, implying that sites located near the coast in early Neolithic times have in all likelihood been lost to the sea.

Timing of the emergence of the Europe–Sicily bridge (40–17 cal ka BP) and its implications for the spread of modern humans

Abstract The submerged sill in the Strait of Messina, which is located today at a minimum depth of 81 m below sea level (bsl), represents the only land connection between Sicily and mainland Italy (and thus Europe) during the last lowstand when the sea level locally stood at about 126 m bsl. Today, the sea crossing to Sicily, although it is less than 4 km at the narrowest point, faces hazardous sea conditions, made famous by the myth of Scylla and Charybdis. Through a multidisciplinary research project, we document the timing and mode of emergence of this land connection during the last 40 kyr. The integrated analysis takes into consideration morphobathymetric and lithological data, and relative sea-level change (both isostatic and tectonic), resulting in the hypothesis that a continental land bridge lasted for at least 500 years between 21.5 and 20 cal ka BP. The emergence may have occurred over an even longer time span if one allows for seafloor erosion by marine currents that have lowered the seabed since the Last Glacial Maximum (LGM). Modelling of palaeotidal velocities shows that sea crossings when sea level was lower than present would have faced even stronger and more hazardous sea currents than today, supporting the hypothesis that earliest human entry into Sicily most probably took place on foot during the period when the sill emerged as dry land. This hypothesis is compared with an analysis of Pleistocene vertebrate faunas in Sicily and mainland Italy, including a new radiocarbon date on bone collagen of an Equus hydruntinus specimen from Grotta di San Teodoro (23–21 cal ka BP), the dispersal abilities of the various animal species involved, particularly their swimming abilities, and the Palaeolithic archaeological record, all of which support the hypothesis of a relatively late land-based colonization of Sicily by Homo sapiens.

Direct radiocarbon dating and genetic analyses on the purported Neanderthal mandible from the Monti Lessini (Italy)

Anatomically modern humans replaced Neanderthals in Europe around 40,000 years ago. The demise of the Neanderthals and the nature of the possible relationship with anatomically modern humans has captured our imagination and stimulated research for more than a century now. Recent chronological studies suggest a possible overlap between Neanderthals and anatomically modern humans of more than 5,000 years. Analyses of ancient genome sequences from both groups have shown that they interbred multiple times, including in Europe. A potential place of interbreeding is the notable Palaeolithic site of Riparo Mezzena in Northern Italy. In order to improve our understanding of prehistoric occupation at Mezzena, we analysed the human mandible and several cranial fragments from the site using radiocarbon dating, ancient DNA, ZooMS and isotope analyses. We also performed a more detailed investigation of the lithic assemblage of layer I. Surprisingly we found that the Riparo Mezzena mandible is not from a Neanderthal but belonged to an anatomically modern human. Furthermore, we found no evidence for the presence of Neanderthal remains among 11 of the 13 cranial and post-cranial fragments re-investigated in this study.